Tuhinga 27: 56–80 Copyright © Museum of New Zealand Te Papa Tongarewa (2016)

A review of the distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand

Sarah E. Jamieson,*‡ Alan J.D. Tennyson,* Kerry-Jayne Wilson,** Elizabeth Crotty,* Colin M. Miskelly,* Graeme A. Taylor*** and Susan M. Waugh* *Museum of New Zealand Te Papa Tongarewa, PO Box 467, Wellington, New Zealand **West Coast Penguin Trust, PO Box 70, Charleston, West Coast, New Zealand ***Department of Conservation, PO Box 10420, Wellington, New Zealand ‡ Current address: Wildlife Research and Monitoring Section, Ministry of Natural Resources and Forestry, c/o Trent University, 2140 East Bank Drive, Peterborough, ON K9L 0G2, Canada

ABSTRACT: Prions are among of the most numerous birds of the Southern Ocean, and yet their populations are poorly documented. New Zealand has breeding populations of four of the six recognised species, all with large population sizes. The remaining two species occur naturally in the New Zealand zone but do not breed there. This review reports data collated from the scientific literature, government archives and unpublished information about the population sizes of prions gathered since earliest scientific records in New Zealand (1773, during James Cook’s second voyage) until the present day. The study focuses on breeding populations, and reports data about population size and presence or absence of prion populations from sites throughout the New Zealand region. The summary presented provides a solid baseline for future population assessments and identifies priority sites where future surveys are warranted.

KEYWORDS: prions, Pachyptila, population sizes, population distribution, Procellariiformes, range, seabirds, New Zealand.

collect the baseline data necessary for robust evaluations of Introduction its population trends and conservation status. Seabirds are one of the most globally threatened groups of Many seabird species rely on remote islands for breeding birds (Croxall et al. 2012). They face a multitude of and nest in burrows, making the collection of even the most pressures, such as interactions with commercial fisheries, basic data challenging. Further, in New Zealand access to pollution, climate change, plastic ingestion and disturbances many of the southern nearshore islands is severely restricted on the breeding grounds, from both humans and invasive to anyone other than traditional owners or occupiers, and species (Carney & Sydeman 1999; Gregory 2009; Hilton & then often limited to the March–May muttonbirding season Cuthbert 2010; Anderson et al. 2011). For many seabird (Moller et al. 2009), a period that does not coincide with the species, adequate knowledge of their distribution is lacking breeding season of many seabird species, including prions. (Croxall et al. 2012). This paucity of the most basic of data Prions (Pachyptila spp.) are small petrels (120–200g inhibits the conservation of these species. Furthermore, a average weight; Miskelly 2013a,b), are nocturnal on poor understanding of a species’ range makes it difficult to land, and nest in burrows or crevices, mostly on remote Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 57

predator-free islands. Globally, there are six species of prion, In this paper we collate data from a wide variety of sources all of which breed on islands in the Southern Ocean: broad- to describe the distribution of prions breeding within the billed prion (P.vittata), Salvin’s prion (P.salvini), Antarctic New Zealand region, and if the data were available, we prion (P.desolata), thin-billed prion (P.belcheri), fairy prion report information on population numbers and trends. (P. t u r t u r ) and fulmar prion (P.crassirostris). Although they From this information we make recommendations for are one of the most abundant groups of seabirds (up to monitoring prion populations with the aim of identifying 95 million individuals; Brooke 2004) and are all listed by colonies that cover the geographic range of each species, the International Union for Conservation of Nature as of but also those that are the most practical to monitor owing ‘Least Concern‘ (IUCN 2016), they still face significant to relative accessibility. Our priority list includes those sites population threats. that have already had some history of monitoring. We also recommend that the population size for each recorded First, prions are among the most common species of colony is estimated. As initial counts of all colonies are seabird to succumb to beach-wreck (e.g. Harper 1980; completed, other priority sites for long-term monitoring Post 2007; Powlesland 1989). For example, during a severe will become apparent. This review does not provide inform - weather event in July 2011, approximately 250,000 prions ation about the biology of prions, nor their distributions (approximately 200,000 of which were broad-billed prions) outside of New Zealand. While we have attempted to blew ashore and died along the west coast of New Zealand include both published and unpublished records to provide (Miskelly 2011a; Tennyson & Miskelly 2011). Climate- a comprehensive overview, it is inevitable that some change models forecast that such storms are likely to increase information will have been missed. in both frequency and intensity (Easterling et al. 2000; Alley et al. 2003). This could have significant detrimental effects on population numbers of prions. Second, prions are Methods surface-feeders that rely on planktonic crustaceans, molluscs A literature review was conducted using primary, secondary and fish. It has been predicted that over the next 90 years and unpublished sources (sources and methods are described there will be a 6.3% decline in ocean productivity (Yool et in Waugh et al. 2013). Raw data were also gleaned from the al. 2013). Much of this decline will be due to a significant authors’ personal field notebooks and those of other decrease in key nutrient levels in surface waters, resulting contributing researchers. We follow the taxonomy and in large-scale effects on the lower trophic levels. In turn, this nomenclature of Gill et al. (2010) and present the results in could resonate throughout the ecosystem (Yool et al. 2013) taxonomic order. and lead to diminished feeding opportunities for surface- We report records of birds on land only, omitting feeding birds. Monitoring even abundant species such as observations of birds on the water, in the air or reported as beach-wrecks. We assumed that the presence of birds ashore prions for assessing changes in marine ecosystems is thus signified breeding; however, birds found in skua middens clearly important; as apex predators, prions are sensitive may have been killed elsewhere (e.g. on the water) and trans- indicators of change throughout these systems. And third, ported to land. Depending on the information available in introduced mammalian predators have extirpated popu - the original source, we described records as individuals (when lations of small seabirds from many islands in New Zealand no information on breeding status was given), breeding pairs since scientific records began 250 years ago (Taylor 2000a). (when some indication of breeding was provided and we Accurate and detailed information about the distribution note the presence of eggs or chicks) or burrows (when we had and abundance of seabirds from the earliest days of scientific information only on the nesting structures themselves, with recording to today would enable these changes to be docu- no information on bird occupancy; note that most prion mented and their impact on species’ conservation status to be nests are in soil burrows, and while in some cases the birds assessed (Warham 1996). However, globally there is a lack of also nest in crevices, such nests are usually also reported as baseline data for most prion populations, with just a few burrows, as most authors did not distinguish between nest exceptions (e.g. Catry et al. 2003; Taylor 2011). The first types). If some level of systematic surveying was conducted, step in determining prion population trends is establishing then the sampling protocol was described as a ‘count‘, the distribution of the species, which is best done during the otherwise it was recorded as an ‘observation‘. A few breeding season, when they are ashore. individual records of live birds on islands well outside their 58 Tuhinga, Number 27 (2016)

50

Observations Counts 40

30

20 Number of records

10

0 Before 1900 1900s 1910s 1920s 1930s 1940s 1950s 1960s 1970s 1980s 1990s 2000s 2010s

Fig.1 Temporal distribution of population records for prion (Pachyptila spp.) colonies within New Zealand. known breeding range (e.g. a broad-billed prion on Motunau 31 records for broad-billed, Antarctic, fairy and fulmar I., Canterbury (Cox et al. 1967) and an Antarctic prion prions, respectively. Prions were found on all major offshore on Houruakopara I., Chatham Is (Imber 1994)) were con- island groups except the Kermadec Is. There were no records sidered to be vagrants. Brief observations that did not add of Salvin’s or thin-billed prions breeding on New Zealand any significant data to more comprehensive observations islands. This was expected, as they are not known to breed were not listed (e.g. there are some records of a species being in the southwest Pacific Ocean (Marchant & Higgins 1990), present on an island when there are other records of actual but it is noted that thin-billed prions were recorded as population estimates from a similar time). possibly breeding at Macquarie I. (Brothers 1984). Fairy We used the names of localities as they are reported by prions had the most expansive New Zealand range, spanning Land Information New Zealand (Land Information New 1650km in distribution from north to south. Antarctic Zealand 2012; Harriss 2016). Island and islands are prions had the most restricted range; they were found almost abbreviated to ‘I.’ and ‘Is‘, respectively. In some cases, text exclusively at the Auckland Is. ‘Observations’ far out- in parentheses after the site name gives information to numbered ‘counts’ (251 versus 51; Fig.1). The number of indicate the location of a small islet, or to reduce ambiguity ‘observations’ peaked during the 1980s, and ‘counts’ peaked about the location, such as where multiple sites with the during the 1990s, after which the number of each decreased same name exist. (likely due to limited resources), but the casual observations The data set associated with this research, including were as numerous as formal counts after the 1990s (Fig.1). detailed latitude and longitude information of the sites, is available to researchers and management groups on request from Susan Waugh at the Museum of New Zealand Te Papa Broad-billed prion Tongarewa (Te Papa; [email protected]). We report broad-billed prions at 48 different locations (Table1). Their colonies spanned from the Chatham Is to the Snares Is/Tini Heke, a distance of c.1400km. Of the 100 Results records for broad-billed prions, only seven colonies had We located 304 records of prion colony observations in the total population estimates based on counts. New Zealand region from the literature and data review; half There were very few repeated observations at any one site of these related to fairy prions. We report 100, 21, 152 and over time, but the few that there were suggested popu lation Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 59

Table1 Population data for broad-billed prions (Pachyptila vittata) nesting in New Zealand (FLD = Fiordland; STW = Stewart I./Rakiura and Foveaux Strait; CIS = Chatham Is; SNI = Snares Is/Tini Heke; dash = no data or comments; see ‘Methods’ for sampling protocol).

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Hawea I., FLD Mar–Apr 10s Burrows Norway rats Observation G. Taylor, Breaksea Sounda 1986 (Rattus norvegicus) unpub. data eradicated Apr 1986

Wairaki I., FLD Mar–Apr 10s Burrows Seals limiting Observation G. Taylor, Breaksea Sounda 1986 nesting sites unpub. data

Gilbert Is FLD Mar–Apr 100s Burrows Dense colony Observation G. Taylor, (western island), 1986 unpub. data Breaksea Sounda

Dusky Sound FLD 1986 – Pairs Breeding Observation K. Morrison in Gaze 1988

Petrel Is, FLD 1785 – Pairs – Observation Begg & Begg Dusky Sound 1968

Anchor I., FLD Mar–May 1000s Pairs – Observation Medway 2011 Dusky Sound 1773 1785 – Individuals Immense numbers Observation Medway 2002 Apr 1900 0 Individuals – Observation Medway 2011

Seal Is, FLD 1773 1000s Pairs – Observation Medway 2011 Dusky Sound

Chalky Inlet FLD 1986 – Pairs Breeding Observation K. Morrison in Gaze 1988

Solander I. FLD Jul 1948 – Burrows Adults Observation Falla 1948 (Hautere) ‘in numbers’; weka patrolling Nov 1973 2 Individuals Corpses Observation Cooper et al. 1986 Feb 1996 100s Pairs Many weka killed Observation A. Tennyson & G. Taylor, unpub. data

Little Solander I. FLD Jul 1985 Several Individuals Seen in flight Observation Cooper et al. 1986

Raratoka I. STW Oct 1989 – Pair 1 chick Observation Cooper 1991 (Centre I.)

Codfish I./ STW Dec 1934 – Pairs Small numbers Observation Wilson 1959: 75 Whenua Hou Dec 1966 – Pairs – Observation Blackburn 1968 2000s 10s Burrows At least 10s of Observation G. Taylor, scattered burrows unpub. data

Sealers Bay stacks, STW 1935 – Pairs – Observation E. Stead in Codfish I./ Blackburn 1968 Whenua Hou Dec 1966 – Pairs – Observation Blackburn 1968

continued on following page 60 Tuhinga, Number 27 (2016)

Table1 Population data for broad-billed prions (Pachyptila vittata) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Sealers Bay stacks, Nov 1991 1000–2000 Burrows – Observation G. Taylor & Codfish I./ A. Tennyson in Whenua Hou [contd ] O’Donnell & West 1998

Trig I. STW Dec 2011 500 Burrows Mostly inactive Count Miskelly 2011b Dec 2011 10 Pairs 10 chicks Count Miskelly 2011b

Green I., STW Nov–Dec 1000s Pairs Many thousands Observation Stead 1953 nr Ruapuke I. 1941 Dec 2012 – Pairs Reported to Observation Miskelly 2013c, be present; unpub. data weka present

Bird I., Mar 1965 – Individuals Large numbers Observation Blackburn 1965 nr Ruapuke I.

North I., STW Oct 1911 – Individuals – Observation Guthrie-Smith 1914 Titi/Muttonbird Is

Jacky Lee I. STW Dec ‘fairly plentiful’ Pairs Many chicks Observation Wilson 1959 (Pukeokaoka) 1932 taken by weka Dec ‘a mere Individuals Decimated Observation Wilson 1959 1940 handful’ by weka

Herekopare I. STW Oct 1911 1000s Individuals – Observation Guthrie-Smith 1914 (Te Marama) May 1942 100s Individuals Cat predation Observation Richdale 1944a observed Dec 1968 0 Individuals – Observation Adams & Cheyne in Fitzgerald & Veitch 1985 Apr–May 1970 1 Individual – Observation Fitzgerald & Veitch 1985

Halfmoon Bay Islet STW 1939/40 16 Individuals Skua midden Observation B. Marples in Anonymous 1953

Whero Rock STW 1941 50 Pairs Count Richdale 1942 1942/43 200 Individuals – Count Richdale 1944a Nov 2010 0 Individuals Nesting site Count Peat 2011 destroyed by shags

Pukeweka I. STW 1931 – Individuals – Observation Wilson 1959

Kundy I. STW Nov 1929 – Pairs – Observation Wilson 1959 Mar 2011 50 Individuals Also 52 in Observation C. Miskelly, skua middens unpub. data

Big I. STW Mar 1965 – Individuals Common Observation Blackburn 1965

Mokiiti/. STW 2006 – Pairs – Observation M. Charteris, Little Moggy I unpub. data

continued on following page Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 61

Table1 Population data for broad-billed prions (Pachyptila vittata) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Mokinui/ STW 2007 – Pairs – Observation M. Charteris, Big Moggy I. unpub. data

Putauhinu I. STW Mar 2011 1 Individual Heard at night Observation C. Miskelly, unpub. data

Tamaitemioka I. STW Mar 1965 – Individuals Skua middens Observation Blackburn 1965

Rerewhakaupoko I. STW Nov 1931 – Pairs – Observation Wilson 1959 (Solomon) Mar 2012 7 Individuals At night Observation C. Miskelly, unpub. data

Pohowaitai I. STW Mar 1965 – Individuals Skua middens Observation Blackburn 1965

Weka I. STW Nov 1931 – Pairs – Observation Wilson 1959

Taukihepa/ STW Jun 1955– – Individuals – Observation Falla in Blackburn Big South Cape I. May 1956 1965

The Sisters CIS Oct 1973 1 Pair 1 chick Observation Imber 1994 (Rangitatahi) Sep 1976 1 Pair 1 egg Observation Imber 1994 (middle island)

S of Owenga, CIS Apr 1983 – Burrows 15 adults killed Observation Imber 1994 Chatham I. by cats

Stack off Cascades CIS Apr 1981 15 Burrows – Count Imber 1994

Blyth’s Stack CIS Nov 1983 18 Pairs Chicks Observation Imber 1994

Houruakopara I. CIS Aug 1980 2 Pairs – Observation Imber 1994 Nov 1987 300 Pairs – Count Plant 1989

Pitt I. (Rangiauria) CIS 1871/72 > 100 Individuals Preyed upon by cats Observation Travers & Travers 1872 1923/24 – Pairs – Observation Archey & Lindsay 1924 1937 – Pairs – Observation Fleming 1939 1951–53 – Pairs Preyed upon by cats Observation Bell 1955 Apr 1967 – Individuals – Observation Imber 1994

Apr 1993 5 Individuals Appeared to be Observation AV36939, killed by cats Canterbury Museum (A. Tennyson, pers. obs.)

Star Keys CIS 1960s–70s 25 Individuals – Observation Imber 1978 Feb 1988 Many Individuals Killed by skuas; Observation A. Tennyson, probably few nesting unpub. data

Rabbit I. CIS Nov 1980 > 100 Pairs – Observation Imber & Lovegrove 1982

continued on following page 62 Tuhinga, Number 27 (2016)

Table1 Population data for broad-billed prions (Pachyptila vittata) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Kokope I. CIS – – – Breeding Observation Imber 1994 Dec 1997 150 Pairs Heavily preyed Observation A. Tennyson, on by weka unpub. data

Mangere I. CIS 1871/72 – Burrows ?Breeding Observation Tennyson & Millener 1994 1923/24 – Burrows – Observation Archey & Lindsay 1924 1937 – Pairs Abundant Observation Fleming 1939 1957 b – Pairs – Observation Tennyson & Millener 1994 1981/82 – Pairs Many large chicks Observation D. Crouchley in Booth 1983 1987/88 10,000 Pairs – Count Tennyson 1989

Little Mangere I. CIS 1937 – Pairs Abundant Observation Fleming 1939 (Tapuaenuku) The Fort

Rangatira CIS Dec 1937 – Pairs Abundant Observation Fleming 1939 (South East I.) Jul 1975 – Individuals Huge numbers Observation Imber 1994 1981/82 – Pairs Many large chicks Observation D. Crouchley in Booth 1983 CIS 1989/90 330,000 Pairs – Count West & Nilsson 1994 1989/90 0.34/m2 Burrows Assumed prion Count West & Nilsson 1994 burrows 1989/90 1.34/m2 Burrows All burrows Count West & Nilsson 1994 Apr 1993 – Individuals Huge numbers Observation Imber 1994 Mar 1999 0.31/m2 Burrows Assumed prion Count Sullivan & Wilson burrows 2001 Apr 2002 1.19±0.10/m2 Burrows All burrows Count Roberts et al. 2007

Western Nugget, CIS Dec 1987 20 Pairs Densely burrowed; Observation Tennyson et al. Murumuru Is partial count 1993

North East I. SNI Jan 1977 – Pair 1 chick Observation Sagar 1977a 1986 2000–5000 Pairs – Observation Miskelly et al. 2001 1986 265 Individuals Skua middens Count Tennyson 2013 Dec 2013 103 Individuals Skua middens Count Tennyson 2013

(South Bay) Feb 1986 350 Individuals – Observation Miskelly et al. 2001 Nov 1986 60 Pairs Chicks; partial count Observation Miskelly et al. 2001 Dec 2013 6 Pairs Chicks; partial count Count Tennyson 2013

Rocky Islet SNI 1971/72 3 Pairs – Observation Horning & Horning 1974 Dec 1976 2 Pairs 2 chicks Observation Sagar 1977a Dec 1984 1 Pair 1 chick Observation Miskelly et al. 2001

continued on following page Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 63

Table1 Population data for broad-billed prions (Pachyptila vittata) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Alert Stack SNI Feb 1985 1 Individual – Observation Miskelly et al. 2001

Broughton I. SNI Nov 1976 – Individuals Killed by skuas Observation P. Sagar in Edgar 1977 Feb 1984 – Individuals – Observation Miskelly et al. 2001

Toru Isletc SNI Dec 1984 2 Pairs 2 chicks Observation Miskelly 1997 Jan 1986 3 Pairs 1 egg, 2 chicks Observation Miskelly et al. 2001

(a) Gaze (1988) and Marchant & Higgins (1990) noted breeding observations from unspecified locations in Breaksea Sound; we provide more detailed observations from this area. (b) Apparently, the year was incorrectly given as 1961 in Tennyson & Millener (1994: table 1), as Lindsay visited in 1957 (Lindsay et al. 1959). However, Lindsay’s diary of this trip, supposedly held in Te Papa’s archives (Tennyson & Millener 1994), could not be located in 2015 (J.Twist, pers. comm.). (c) Note that while broad-billed prions have been reported from skua middens on Rima Islet (Snares Is/Tini Heke; Sagar 1977b), there is no evidence that the species breeds there (C.Miskelly, unpub. data). declines. For example, thousands of birds were believed to Sealers Bay stacks off Codfish I./Whenua Hou (1000–2000 have been nesting on Anchor I. in Dusky Sound, Fiordland, burrows in 1991; Table 1). However, the number of birds during James Cook’s visit in 1773; however, by 1900 a in the Fiordland and Stewart I./Rakiura regions is poorly breeding population was no longer present there (Medway known and more detailed surveys are warranted there. 2011). Similar declines resulting in localised extinction are suspected at the colony on the neighbouring Seal Is Antarctic prion (Medway 2011). To the east on Kokope I. (Chatham Is), and The Auckland Is are the stronghold of Antarctic prions at Solander I. (Hautere) and Jacky Lee I. (Pukeokaoka) breeding in New Zealand (Table2); the species has been (both in the Stewart I./Rakiura region), the populations reported from eight different islands in the group. However, were being heavily depredated by weka (Gallirallus australis). there are no substantive data from which to assess Prions trying to nest on Chatham I. were reported to be population size among the 19 records for the species in the heavily preyed upon by cats (Felis catus; Imber 1994). On group, and so we are unable to draw conclusions about their Herekopare I. (Te Marama), off Stewart I./Rakiura, the population status or trends. population went from thousands of individuals in 1911 to a single individual in 1970, reportedly due to cat preda tion (Guthrie-Smith 1914; Fitzgerald & Veitch 1985). The entire Fairy prion population on Whero Rock was extirpated after colonisa- With numbers of breeding pairs in the millions, fairy prions tion by the New Zealand endemic Stewart Island shag are among the most common seabird species nesting in (Leucocarbo chalconotus), whose nesting activity destroyed New Zealand (sooty shearwater, Puffinus griseus, is the most the small cap of vegetation on the islet (Richdale 1944a; abundant; see Waugh et al. 2013 for a population assess - Peat 2011). The Chatham Is host the largest portion of ment). They are also one of the most widespread New New Zealand’s breeding population of broad-billed prions, Zealand seabird species, with a geographic range extending yet there have not been any repeated counts at the large from the Poor Knights Is in Northland to Stewart I./Rakiura colony at Mangere I. and only a limited number of counts and the Antipodes Is. (Table3). The largest colony within conducted at Rangatira (South East I.), the most recent New Zealand, and likely the world, is on Stephens I./ being in 2002. Despite huge colonies of prions formerly Takapourewa in the Marlborough Sounds, which has about being present in southern New Zealand, the largest known 1.4 million pairs (Craig 2010). The limited information documented remaining colonies in recent times are at the available suggests that the second-largest colony in the New Snares Is/Tini Heke (2000–5000 pairs in 1986) and on the Zealand region is on Mangere I. in the Chatham Is, with 64 Tuhinga, Number 27 (2016)

Table2 Population data for Antarctic prions (Pachyptila desolata) nesting in New Zealand (AKI = Auckland Is; CBL = Campbell I./Motu Ihupuku; dash = no data or comments; see ‘Methods’ for sampling protocol).

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Auckland Isa AKI 1984 100,000– Pairs – Unknown Robertson & Bell 1,000,000 1984 1990 350,000– Pairs – Unknown Marchant & Higgins 750,000 1990

Enderby I. AKI 1944 – Pairs Common Observation Turbott 2002 Jan 1966 – Burrows – Observation Taylor 1971 Dec 1976 – Individuals Skua middens Observation Bartle & Paulin 1986 Feb 1988 – Pairs Calling from Observation G. Taylor, burrows, 1 adult unpub. data seen in burrow

Rose I. AKI Jan 1966 – Burrows – Observation Taylor 1971

Auckland I. AKI 1907 – Burrows Commonb Observation Waite 1909 1944 – Pairs Common Observation Turbott 2002 Feb 1973 – Pairs Most common petrel Observation Challies 1975 Feb–Mar 100+ Individuals Killed by cats Observation Thompson 1986 1982 Feb 1988 – Individuals Many birds Observation G. Taylor, killed by cats unpub. data outside burrows

Ocean I. AKI 1972 4 Individuals – Observation K.-J. Wilson, unpub. data Feb 1988 – Pairs Calling from Observation G. Taylor, burrows unpub. data

Shoe I. AKI 1903 – Burrows – Observation Waite 1909

Disappointment I. AKI Nov 1907 – Individuals Remains only Observation Waite 1909

Adams I. AKI 1944 – Pairs Common Observation Turbott 2002 Nov 1989 – Pairs – Observation Buckingham et al. 1991

Masked I. AKI Nov 2013 2 Pairs – Count K.-J. Wilson, unpub. data

Northwest Bay CBL Jan 1986 1 Individual Found in a burrow; Observation D. Cunningham, stack probably this species pers. comm. to G. Taylor

Eboulé Peninsula, CBL Jan 2006 1 Individual Fledgling in Observation Miskelly & Campbell I./ skua midden; Fraser 2006 Motu Ihupuku not clear evidence of breeding at this site

(a) Te Papa holds specimens of Antarctic prions that indicate additional or probable breeding islands in the Auckland Islands group: 1 egg (NMNZ OR.14749, collected 4 Dec 1943) and 1 chick (NMNZ OR.13031, collected 14 Jan 1943) from Figure of Eight Island; 2 adults (NMNZ OR.17547 and OR.17548, both collected 8 Jan 1973) from Ewing Island; 1 complete skeleton (NMNZ OR.19794, collected 21 Feb 1973) from Monument Island. (b) Waite (1909) was uncertain if the burrows were created by Antarctic or broad-billed prions; the latter have never been observed in AKI, so we have assumed the burrows were made by Antarctic prions. Also, Antarctic prions were reported by Waite (1909) as being on the Antipodes Is but likely a case of misidentification, as only fairy prions have been sighted there by other observers (Tennyson et al. 2002). Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 65

Table3 Population data for fairy prions (Pachyptila turtur) nesting in New Zealand (NL = Northland; KAP = Kapiti coast; MLS = Marlborough Sounds; WCN = west coast, North I.; WCS = west coast, South I.; CTC = Canterbury coastal; OTC = Otago coastal; FLD = Fiordland; STW = Stewart I./Rakiura; CIS = Chatham Is; SNI = Snares Is/Tini Heke; ANT = Antipodes Is; dash = no data or comments; see ‘Methods’ for sampling protocol).

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Poor Knights Is NL 1930s – Pairs Burrows and eggs Observation Falla 1934

Tawhiti Rahi I., NL Jan 1943 – Pairs Small numbers Observation Buddle 1946 Poor Knights Is Aug 1958 – Individuals Moderate numbers Observation Kinsky & Sibson 1959 Dec 1958 1 Pair Downy chick Observation Kinsky & Sibson 1959 Sep 1980 1000s Pairs Many thousands Observation McCallum 1981

Aorangi I., NL Nov 1940 – Pairs Vast numbers Observation Buddle 1941 Poor Knights Is Aug 1958 – Individuals Moderate numbers Observation Kinsky & Sibson 1959 1964–75 40,000 Individuals Extrapolation Count Harper 1976 from plot surveys Nov 1990 – Pairs Many; some eggs Observation R. Parrish in Taylor & Parrish 1992 Dec 2011 – Pairs Widespread Observation G. Taylor & A. Tennyson, unpub. data

Te Haupa I. NL Prior to 1934 – Pairs Chicks Observation Falla 1934 (Saddle I.), Apr 1990 0 Individuals Ship rats present Observation G. Taylor & off Great Barrier I. A. Tennyson, (Aotea I.) unpub. data Nov 1994 0 Individuals Ship rats present Observation A. Tennyson & K. McConkey, unpub. data

Hauturu/ NL 1886 – Individuals – Observation Reischek 1887 Little Barrier I. Nov 1960 1 Individual Presumed vagrant Observation Bishop 1963 Dec 1962 1 Individual Presumed vagrant Observation Bishop 1963 1978–2015 0 – No recent sightings Observation A. Tennyson, G. Taylor & C. Miskelly, unpub. data

Mana I. KAP 2005 1 Nest – Count Miskelly & Gummer 2013 2008 3 Pairs – Count Miskelly 2010 2012 6 Pairs – Count Miskelly & Gummer 2013

Stephens I. MLS 1925 100,000s Burrows – Observation Guthrie-Smith 1936 (Takapourewa) May 1974– 0.2– Burrows Range Count Walls 1978 Apr 1975 4.5/m2 1985 1,000,000 Individuals – Unknown Harper 1985 1990 500,000 Individuals – Unknown Daugherty et al. 1990 Jun–Dec 0.5±0.3– Burrows Varied by habitat Count Markwell 1997 1994 1.4±0.2/m2 (means ± SEs)

continued on following page 66 Tuhinga, Number 27 (2016)

Table3 Population data for fairy prions (Pachyptila turtur) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Stephens I. Jun–Dec 0.4±0.1– Pairs Varied by habitat Count Markwell 1997 (Takapourewa) [contd ] 1994 1.1±0.2/m2 (means ± SEs) Aug 1994 0.095/m2 Burrows Mean (range) Count Craig 2010 (0–0.371/m2) Aug 1994 1,830,523 Burrows – Count Craig 2010 Aug 1994 1,418,665a Pairs Occupancy rate Count Craig 2010 0.775 (Craig 2010: table 14) 1998 0.84/m2 Burrows Mean (range) Count Mulder & Keall 2001 (0–3/m2)

Jag Rocks MLS 1961 – Individuals – Observation B. Bell, unpub. data Apr 1987 – Burrows Abundant Observation G. Taylor, unpub. data

Middle Trio I., MLS Apr 1963– – Pairs Numerous Observation Campbell 1967 Trio Is Jan 1964 (Kuru Pongi) 1990 – Individuals – Observation Daugherty et al. 1990

Sentinel Rock MLS Apr 1987 – Individuals Feathers common Observation G. Taylor, in crevices unpub. data

Ninepin Rock, MLS Aug 1993 – Burrows Numerous Observation D. Brown in nr Chetwode Is O’Donnell 1995

The Haystack MLS Aug 1993 – Burrows Numerous Observation D. Brown in (Moturaka), O’Donnell 1995 nr Chetwode Is

North Brother I. MLS Aug 1950– – Pairs In great numbers Observation Sutherland 1951 Feb 1951 Oct 1990 1000 Pairs – Count Gaston & Scofield 1995 Oct 1990 0.03/m2 Pairs Mean (range) Count Gaston & Scofield (0–5/m2) 1995 Oct 1990 1.4/m2 Burrows Mean (range) Count Gaston & Scofield (0–14/m2) 1995 Feb 1993 1750 Burrows Medium reliability Count K.-J. Wilson, unpub. data

South Brother I. MLS Early 1960s – Individuals – Observation B. Bell & I. Crook, unpub. data

The Twins MLS 1961 – Individuals – Observation B. Bell, unpub. data

Motungarara I. MLS 1961 – Individuals – Observation B. Bell, unpub. data

Wall I. WCS Dec 2013 1255 Pairs – Count R Lane & M. Charteris in R. Lane, unpub. data Dec 2015 1400 Pairs – Count R. Lane, unpub. data

continued on following page Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 67

Table3 Population data for fairy prions (Pachyptila turtur) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Motukiekie Rocks WCS 2000 – Burrows – Observation G. Wood, unpub. data (islet) Jan 1995 8 Nests 8 chicks Observation B. Stuart-Menteath in O’Donnell & West 1996

Murphy Beach WCS Mar 2010 2 Nests 1 egg, 1 chick Observation OR.029176 and stacks OR.029213, Te Papa

Arnott Point islet WCS Mar 2010 1 Nest 1 chick Observation OR.029177, Te Papa

Taumaka I., WCS 1907 – Nests See note b Observation Waite 1909 Open Bay Is Feb 1973 – Nests Uncommon Observation K.-J. Wilson, unpub. data Oct 1980 – Nests Uncommon Observation K.-J. Wilson, unpub. data Aug 1986 10s Nests Some weka Observation A. Tennyson, predation unpub. data 1994/95 – Nests Chick remains Observation Miller 1997

Barn Is WCS Mar 2011 – Burrows Numerous Observation Lettink et al. 2013

Motunau I. CTC 1958 9900 Burrows – Observation Cox et al. 1967 1962 14,000 Burrows – Count Cox et al. 1967 1961–63 27,500 Individuals Very rough Count Cox et al. 1967 estimate Dec 1983 – Pairs Many Observation J. Fennell & P. Sagar in Gaze 1985 Dec 1996– 14,000 Burrows – Count Beach et al. 1997 Jan 1997 2004 – Individuals Harrier midden Observation Hawke et al. 2005

Crown I. CTC 1960 – Pairs Nesting densely Observation B. Bell in Wilson 2008c (Le Bons Bay to Dec 2000 255 Burrows – Count Wilson 2008 Pompeys Pillar)

Islet, CTC Dec 2000 30 Pairs – Count Wilson 2008 Redcliffe Nook

Islet, Island Nook CTC 1960 – Pairs – Observation B. Bell in Wilson 2008 Dec 2000 150 Pairs – Count Wilson 2008

Islet, Island Bay CTC 1960 – Pairs – Observation B. Bell in Wilson 2008 Dec 2000 300 Pairs – Count Wilson 2008

Wharekakahu OTC Nov 1983 2000–3000 Burrows Occupancy rate Count Ward & Munro 1989 70% at plot examined

Gull Rocks OTC 1990s – Pairs – Observation Loh 2000

continued on following page 68 Tuhinga, Number 27 (2016)

Table3 Population data for fairy prions (Pachyptila turtur) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Green I. OTC 1980s – Pairs Important breeding area Observation Ward & Munro 1989 No date – Pairs – Observation Loh 2000

Tunnel Beach OTC Feb 1997 8 Burrows – Count Loh 2000 (Prion Cave) Aug 1996 – Individuals – Observation Loh 2000 (Prion Cleft) Jan 1993 70 Individuals 14 potential nests Count Loh 2000 (Prion Cliff) Oct 1998 160 Individuals Artificial nests Count Loh 2000 installed from 1994

Rock stacks, Catlins OTC No date – Pairs ‘Small colonies’ Observation Loh 2000

Solander I. FLD Jul 1948 – Burrows Weka patrolling Observation Falla 1948 (Hautere) Jan 1973 – Individuals Bones Observation Wilson 1973 Nov 1973 – Pairs Small colony Observation Cooper et al. 1986 Nov 1976 – Individuals Small numbers Observation Cooper et al. 1986 Feb 1996 100s Pairs Many killed Observation A. Tennyson & by weka G. Taylor, unpub. data Jul 1997 – Pairs Scattered in areas Observation G. Taylor, inaccessible to weka unpub. data

Little Solander I. FLD Jul 1948 – – Remains in ‘skua and Observation Falla 1948 (or) hawk castings’ Nov 1976 – Individuals Small numbers Observation Cooper et al. 1986 Jul 1985 1 Individual Seen in flight Observation Cooper et al. 1986

Codfish I./ STW Dec 1934 – Pairs Small numbers Observation Wilson 1959: 75 Whenua Hou 1991–2011 0 – No recent sightings Observation G. Taylor, A. Tennyson & C. Miskelly, unpub. data

Green I. STW Nov 1941 1,000,000 Burrows – Count Stead 1953 (<1/m2) Nov 1941 1,500,000d Pairs – Count Wilson 1959 Dec 2012 – Individuals Only 6 corpses; Observation Miskelly 2013c, weka present unpub. data

North I., Titi/ STW Oct 1911 – Individuals – Observation Guthrie-Smith 1914 Muttonbird Is

Jacky Lee I. STW Dec 1932 – Pairs ‘Fairly plentiful’; Observation Wilson 1959 (Pukeokaoka) many taken by weka Dec 1940 – Individuals ‘A mere handful’; Observation Wilson 1959 decimated by weka

Herekopare I. STW Oct 1911 1000s Individuals – Observation Guthrie-Smith 1914 (Te Marama) Aug 1941 10s Individuals Some dozens Observation Richdale 1944b Dec 1968 – Pairs A very large Observation Adams & Cheyne population in Fitzgerald & Veitch 1985

continued on following page Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 69

Table3 Population data for fairy prions (Pachyptila turtur) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Halfmoon Bay Islet STW 1939/40 44 Individuals Skua midden Observation Anonymous 1953

Bench I. STW Nov 1971 1 Individual – Observation K.-J. Wilson, unpub. data

Whero I. STW 1940s 1000 Individuals – Count Richdale 1965 1941 400 Pairs – Count Richdale 1942 2010 0 Individuals – Count Peat 2011

Kundy I. STW Nov 1929 – Pairs – Observation Wilson 1959 Mar 2011 1000s Individuals Also 44 in Observation C. Miskelly, skua middens unpub. data

Mokiiti/ STW 2007 – Pairs NW and NE Observation M. Charteris, Little Moggy I. headlands unpub. data

Big I. STW Mar 1965 – Individuals Carcasses Observation Blackburn 1965 Kaimohu I. STW Feb 1965 – Individuals Skua middens Observation Blackburn 1965

Putauhinu I. STW Mar 2011 1 Individual Heard at night Observation C. Miskelly, unpub. data

Tamaitemioka I. STW Mar 1965 – Individuals Skua middens Observation Blackburn 1965 Rerewhakaupoko I. STW Nov 1931 – Pairs – Observation Wilson 1959 (Solomon) Jan 1955– – Individuals – Observation Falla in Blackburn May 1956 1965

Pohowaitai I. STW Dec 1929 – Pairs In burrows Observation E. Stead diary (C. Miskelly, unpub. data) Mar 1965 – Individuals Skua middens Observation Blackburn 1965

Taukihepa/ STW Jun 1955– – Individuals – Observation Falla in Blackburn Big South Cape I. May 1956 1965 Apr 1961 – Individuals Common Observation Bell & Merton in Blackburn 1965 Aug 1964 – Individuals – Observation Bell & party in Blackburn 1965

Chatham Is CIS 1871/72 – Pairs ‘Immense numbers’ Observation Travers & Travers 1872

The Sisters CIS No date – Pairs – Observation Imber 1994 (Rangitatahi) (western island) (middle island) Jan 1954 2 Pairs – Observation Dawson 1955; Marchant & Higgins 1990 Jan 1974 1 Nest 1 chick Observation Imber 1994

Star Keys CIS 1960s–70s 25 Individuals – Observation Imber 1978 Feb 1988 Many Individuals Killed by skuas; Observation A. Tennyson, probably few nesting unpub. data

continued on following page 70 Tuhinga, Number 27 (2016)

Table3 Population data for fairy prions (Pachyptila turtur) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Rabbit I. CIS Oct–Nov 100s Individuals Eggs noted Observation Imber & Lovegrove 1980 1982; Imber 1994 Apr 1981 – Individuals Many visiting Observation Imber 1994

Kokope I. CIS No date – Pairs – Observation Imber 1994 Dec 1997 0 Individuals – Observation A. Tennyson, unpub. data

Mangere I. CIS 1871/72 – Individuals Probably in Observation Travers & Travers ‘immense numbers’ 1872 1923/24 – Burrows Numerous Observation Archey & Lindsay 1924 1937 – Pairs – Observation Fleming 1939 Jul 1975 – Individuals Abundant Observation Imber 1994 Oct 1980 1000s Pairs Some thousands Observation T.G. Lovegrove in Booth 1982 1987/88 30,000 Pairs – Count Tennyson 1989e

Little Mangere I. CIS 1937 – Pairs – Observation Fleming 1939 (Tapuaenuku) The Fort

Western Nugget, CIS Dec 1987 1 Nest Partial count Observation Tennyson et al. 1993 Murumuru Is

Daption Rocks SNI Feb 1977 – Individuals Killed by skuas Observation Miskelly et al. 2001 (north) (south) Nov 1976 – Pairs – Observation Miskelly et al. 2001

North East I. SNI 1985–87 3500 Pairs – Count Miskelly et al. 2001

Rocky Islet SNI Dec 1971 3 Individuals – Observation K.-J. Wilson, unpub. data Dec 1976 – Pairs – Observation Miskelly et al. 2001 Dec 1984 3 Pairs 3 eggs Observation Miskelly et al. 2001

Alert Stack SNI Dec 1976 – Pairs – Observation Miskelly et al. 2001

Broughton I. SNI Feb 1984 – Individuals – Observation Miskelly et al. 2001 Mar 1992 500 Pairs – Observation Miskelly et al. 2001

Antipodes Is ANT Jan–Mar 0 Individuals Unable to find Observation Warham & Bell 1979 (all islands) 1969 any on land Apr 2001 – Individuals Scores Observation Imber et al. 2005

Antipodes I. ANT Nov–Dec 20+ Pairs Not in large Observation Imber 1979, 1983 1978 numbers Nov 1995 1000–5000 Pairs – Observation Tennyson et al. 2002

Bollons I. ANT Nov–Dec – Pairs – Observation Imber 1979, 1983 1978

continued on following page Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 71

Table3 Population data for fairy prions (Pachyptila turtur) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Archway I. ANT Nov–Dec – Individuals – Observation Imber 1979 1978

(a) This value differs from the population total of 2,160,017 pairs in Craig (2010). Craig’s total assumes that burrow count equals nesting population, but values from her table 14 suggest a burrow occupancy rate of 0.775, and hence a population of 1,418,665 breeding pairs (1,850,523 × 0.775) is more accurate. (b) Listed as probably Antarctic prions but assumed to be fairy prions as they are the only species to have been recorded nesting in the area by other observers. (c) A fledgling was collected by E. Stead and A. Brooks Jr. in 1935 at an ‘Islet near Akaroa Inlet’ (Museum of Vertebrate Zoology, Berkeley (MVZ Birds 72373), retrieved on 9 April 2014 from http://arctos.database.museum/guid/MVZ:Bird:72373) – this location is probably one of the islets discussed by Wilson (2008). (d) Assumed to be primarily fairy prions, although there are likely to be some broad-billed prions included in this count as Stead (1953) estimated a ratio of one broad-billed prion to every 12 fairy prions. (e) An estimate of 40,000 pairs (Taylor 2000b; Aikman & Miskelly 2004) is based on a misquotation of Tennyson (1989), which states 30,000 pairs.

30,000 pairs (Tennyson 1989), followed by Aorangi I. in the presumably happening as a result of land being retired from Poor Knights Is, with 40,000 individuals (Harper 1976), farming and the habitat improvements associated with exten- then Motunau I. in Canterbury, with 14,000 burrows (Cox sive planting of new forest areas (currently coordinated by et al. 1967), although admittedly some of these data are the Department of Conservation). Stock trampling of bur- decades old and the current sizes of these colonies may rows in the past would have reduced burrow densities over have changed significantly. More than 2000 pairs are also large parts of the island (Taylor, pers. obs.). likely to nest on Tawhiti Rahi I. in the Poor Knights Is Fairy prion numbers appear to have remained stable at (McCallum 1981), Wharekakahu in Otago (Ward & Wall I. in Westland, with between 1255 and 1400 pairs Munro 1989), Herekopare I. (Te Marama) and Kundy I., recorded in 2013 and 2015 (R.Lane, unpub. data). The tiny both in the Stewart I./Rakiura region (Guthrie-Smith 1914; Mana I. colony has been slowly growing after it was C.Miskelly, unpub. data), North East I. in the Snares Is/ established using translocated chicks (Miskelly & Gummer Tini Heke (Miskelly et al. 2001) and Antipodes I. (Tennyson 2013). The Motunau I. population appears to have et al. 2002). remained stable from 1962 to 1996 (Cox et al. 1967; Beach Although the species was reported in 66 locations from et al. 1997). In contrast, six other colonies appear to have 152 records, population trends can be estimated at only nine decreased in size or been extirpated. colonies. The largest apparent increase was at Stephens I. The most substantial decline occurred on Green I., (Takapourewa), where reported numbers went from 1mil- northeast of Stewart I./Rakiura. An estimated 1.5 million lion individuals in 1985 to 1.4million pairs in 1994 (Harper pairs were nesting on the island in November 1941 (Wilson 1985; Craig 2010; this study). A five- to sixfold increase in 1959), yet in December 2012 there was very little sign of any population size over nine years is highly improbable, so these nesting prions (Miskelly 2013c). Weka are present on the differences presumably reflect differing methodologies. island (Miskelly 2013c), but it is unclear if weka predation However, a more detailed examination of data reported by could have caused such a massive decline in prion numbers. Walls (1978) and Mulder & Keall (2001) does indicate that On nearby Jacky Lee I. (Pukeokaoka), where weka were the population on Stephens I. (Takapourewa) has grown not harvested by muttonbirders, the dense weka population over time. Both studies conducted surveys in the same area all but extirpated fairy prions within a few decades (Wilson of the island (Keeper’s Bush), and found that the density of 1959). On neighbouring Whero I., the population declined burrows increased from 0.70/m2 in 1975 to 0.95/m2 in 1998 from 1000 individuals in the 1940s to a total absence in (note that the numbers reported in Table 3 are for the entire 2010 after an expanding colony of Stewart Island shags island, not just Keeper’s Bush). This population increase is destroyed the vegetation on the islet (Richdale 1965; Peat 72 Tuhinga, Number 27 (2016)

Table4 Population data for fulmar prions (Pachyptila crassirostris) nesting in New Zealand (CIS = Chatham Is; BIS = Bounty Is; SNI = Snares Is/Tini Heke; AKI = Auckland Is; dash = no data or comments; see ‘Methods’ for sampling protocol).

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Chatham Is CIS 1984 1000– 5000 Pairs – Unknown Robertson & Bell (total for all islands) 1984

Motuhara CIS 1974 – Individuals – Observation Imber 1994 (Forty-Fours) Dec 1983 – Pairs Many Observation Imber 1994

The Pyramid CIS Dec 1937 – Pairs Numerous carcasses, Observation Fleming 1939 (Tarakoikoia) 2 nests Nov 1974 – Pairs – Observation Imber 1994 Dec 1987 7 Pairs Partial count Observation Tennyson et al. 1993

Bounty Is BIS 1888 – Nests See note a Observation A. Reischek in (total for all islands) Robertson & van Tets 1982 1907 – Nests See note a Observation Waite 1909 Nov 1978 76,000 Pairs ‘Impossible to census’ Observation Robertson & van Tets 1982 1997 29,354 Pairs Medium reliability Count Booth & Schmechel in Taylor 2000a 1997 0.21/m2 Nests Medium reliability Count Booth in Taylor 2000a

Proclamation I. BIS Dec 1997 1235 Pairs Good reliability Count Booth in Taylor 2000a

Toru Islet SNI Dec 1972 – Pairs – Observation Fleming & Baker 1973 Nov 1976 – Pairs – Observation Sagar 1977b

continued on following page

2011). In 1934, a small fairy prion colony was apparently in Northland in the nineteenth century was probably present on Codfish I./Whenua Hou (Wilson 1959), but misreported or a mis - identification of another petrel species weka and Pacific rats (Rattus exulans) were also present and (see Falla 1934; Skegg 1964). the species has not been reported since, despite extensive Breeding fairy prions were reported on Kokope I. in the research and management work at the site. Chatham Is by Imber (1994), but the species was not found In 1886, fairy prions were reported to be present ashore in 1997 and there was considerable evidence of on Hauturu/Little Barrier I. (Reischek 1887), but again predation on other petrel species by weka (A.Tennyson, they have not been reported since. Cats and Pacific rats unpub. data). were present, so the colony may have been extirpated by them. The population on Te Haupa (Saddle I.), off Fulmar prion Great Barrier I. (Aotea I.) appears to have been extirpated Fulmar prion colonies were reported on the Chatham Is, (reportedly by ship rats, Rattus rattus), although there are Bounty Is, Snares Is/Tini Heke and Auckland Is (Table4), no data on the initial size of the population (Falla 1934). but from only eight individual islands within these groups, This suggests that the geographically isolated population and with counts from just two sites of the total 31 records. currently nesting on the Poor Knights Is is a remnant The Bounty Is are home to the largest population of of a more widespread northern New Zealand population. breeding birds (29,354 pairs in 1997; Booth & Schmechel A fairy prion colony supposedly on the Marotere Is in Taylor 2000a). This was a significant decline from the Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 73

Table4 Population data for fulmar prions (Pachyptila crassirostris) nesting in New Zealand. Continued from previous page

Locality name Area Dates Counts Status Occurrence Sampling Reference comments protocol

Toru Islet [contd ] Feb 1984 300–400 Pairs Later doubted Observation Miskelly 1984; by author Miskelly et al. 2001 Dec 1984 4 Pairs 2 eggs, 2 chicks Observation Miskelly et al. 2001 Jan 1986 6 Pairs 2–6 eggs and chicks Observation Miskelly et al. 2001 Sep–Oct 2010 – Individuals Low numbers, Observation Carroll & Charteris widely distributed 2010 Nov 2013 100s Pairs – Observation A. Tennyson & C. Miskelly, unpub. data

Rima Islet SNI Nov 1976 – Pairs – Observation Sagar 1977b Feb 1984 100–200 Pairs – Observation Miskelly 1984 Sep 2010 – Individuals Low numbers, Observation Carroll & Charteris widely distributed 2010

Auckland Is AKI 1984 1000–5000 Pairs – Unknown Robertson & Bell (total for all islands) 1984 1998 <1000 Pairs – Observation Tennyson & Bartle 2005

Rose I. AKI 1943 – Pairs Small numbers Observation Taylor 1971 Nov 1972– – Individuals – Observation Bell 1975 Mar 1973 1998 Few 100 Pairs Estimate based on Observation Tennyson & Bartle densities nearby 2005

Ocean I. AKI Jun 1998 <100 Pairs – Observation Tennyson & Bartle 2005

Ewing I. AKI Nov 1972– – Individuals – Observation Bell 1975 Mar 1973 Nov 1989 – Individuals – Observation Moore & McClelland 1990 Jun 1998 100–400 Pairs – Observation Tennyson & Bartle 2005

(a) Listed as Pachyptila turtur [= fairy prion] in Robertson & van Tets (1982) and Prion desolatus [=Antarctic prion] in Waite (1909) but assumed to be fulmar prions as they are the only species to have been recorded nesting at this island group by other observers.

1978 population estimate of 76,000 pairs, although as the estimate (and no information was given about the sampling researchers described them as ‘impossible to census’ protocol used), this difference cannot be considered real, and (Robertson & van Tets 1982) it is difficult to be certain clearly there is a need for new and accurate surveys of the that the apparent decline is real. Apart from the colony at prion populations for this island group. Proclamation I., the distribution of nesting islands within the Bounty Is group has not yet been reported. The population nesting on the Auckland Is in 1984 was Discussion estimated to be 1000–5000 pairs, but less than 1000 pairs We located 304 records of prions breeding on islands 14 years later (Robertson & Bell 1984; Tennyson & Bartle throughout New Zealand and its subantarctic and offshore 2005). Because such a wide range was reported for the 1984 islands, except the Kermadec Is. There are no population 74 Tuhinga, Number 27 (2016)

Table5 Population sizes and trends, and quality of information for four species of prion (Pachyptila spp.) nesting in the New Zealand region.

Total population Estimated number of breeding sites Trend Quality of Species estimate (number of sites with population information information (breeding pairs) estimates or counts since 1995)

Broad-billed prion 350,000 44 (6) Unknown Poor

Antarctic prion 350,000–1,000,000 8 (0) Unknown Poor

Fairy prion 1,500,000 64 (1 ) Unknown Poor

Fulmar prion 31,000–36,000 8 (4) Unknown Poor

estimates or counts for the majority of prion breeding sites causes of decline are not always obvious. For instance, and the data consist of five times more ‘observations’ than Rangatira (South East I.) is free of introduced predators ‘counts’. Moreover, there were very few sites where repeat (Aikman & Miskelly 2004), yet the density of broad-billed counts have been conducted. For fairy and broad-billed prion burrows between 1989 (0.34/m2) and 1999 (0.31/m2) prions, only 17% and 14% of colonies, respectively, have has decreased by 8% and the rate of decline is even steeper if had their breeding populations estimated within the last burrows of all sizes (1.34/m2 in 1989, 1.19/m2 in 2002) are 20 years (Table5). No Antarctic prion colony estimates exist included (11%). There are no obvious reasons for this other than broad overall population estimates from more change. Western Gilbert I. (Fiordland), Sealers Bay stacks than 25 years ago, with no reference to how these numbers and Trig I. (Stewart I./Rakiura region), Mangere I. and were attained. Rangatira (South East I.) (Chatham Is), and North East I. This lack of data severely limited our ability to draw any (Snares Is/Tini Heke) may be good candidates for establish- conclusions about population sizes or to assess population ing regular long-term survey plots. They are widely dispersed trends. The paucity of information emphasises the need to throughout the broad-billed prion’s New Zealand range. collect baseline data. Without reliable information on Furthermore, each site already has some level of baseline population size and trend, decisions about management of data and most are regularly visited by seabird researchers. threats are problematic. It is concerning that the number of Due to the lack of data, it is impossible to estimate the population records has decreased since the 1980s–1990s, current size of the New Zealand’s breeding population of but offsetting this is the fact there has been an increase in the Antarctic prions. Estimates as high as 750,000 (Harper in accuracy and repeatability of surveys since the 1980s. Marchant & Higgins 1990) and 1million pairs (Robertson There was evidence of population declines at several of the & Bell 1984) have been published, but neither of these few broad-billed prion colonies where data allowed us to accounts provides any information about how the figures assess population trends. Population declines were due to were determined. Within the New Zealand region, Antarctic both predation and habitat destruction by other seabirds. prions have been confirmed nesting only on the Auckland For example, the population on Anchor I. was likely to have Is. In the nineteenth century, these islands saw the arrival of been extirpated by stoats (Mustela erminea; see Medway sealers and whalers. This led to the introduction of mammals 2011). On Herekopare I. (Te Marama), cats extirpated the such as European rabbits (Oryctolagus cuniculus), cats, house broad-billed prion population (Fitzgerald & Veitch 1985), mice (Mus musculus) and pigs (Sus scrofa) (Taylor 1971), and cats continue to cause significant damage to wildlife which today remain predators of ground-nesting birds or populations at other sites, such as Chatham I. (Imber 1994). cause significant habitat destruction. Some islands in this The populations of both broad-billed and fairy prions on group (including Adams I. and Disappointment I.) have Whero Rock were extirpated due to habitat destruction remained free of introduced mammals. Enderby I. and Rose caused by Stewart Island shags (Peat 2011). However, the I. had diverse introduced mammal communities, but these Distribution and size of prion (Pachyptila spp.) colonies throughout New Zealand 75

mammal species were eradicated in the early 1990s (Torr nesting on Australia’s Heard I. nesting in New Zealand 2002). Auckland I. is the most heavily impacted by (Tennyson & Bartle 2005). However, our analysis suggests introduced mammals of all the islands in the group, and that the global population estimate is likely to be too high: is the only island still supporting introduced mammals the Bounty Is, where most fulmar prions breed, are estimat- (pig, cat and house mouse) (Taylor 1968; Taylor 2000a). ed to have a nesting population of approximately 30,000 Monitoring Antarctic prions on Enderby I. and Adams I. is pairs (Taylor 2000a). The only other colonies of significant considered a high priority, particularly as these sites are size are in the Chatham Is and Auckland Is, both of which regularly visited by researchers, while at Ewing I. the recent Robertson & Bell (1984) estimated at 1000–5000 pairs. expansion of the Olearia forest (K.-J.Wilson, pers. obs.) Robertson & Bell (1984) gave no information about how indicates changes occurring at the site, which have potential these numbers were arrived at, and more recent work by to impact on the prion numbers. Tennyson & Bartle (2005) suggests that the Auckland Is Fairy prions are the most numerous and widespread population is less than 1000 pairs. The remaining site where species of prion nesting in New Zealand. The largest the species is known to breed is the Western Chain of the population, on Stephens I. (Takapourewa), numbers Snares Is/Tini Heke, which is thought to hold less than 1000 approximately 1.4 million pairs and appears to be growing pairs, resulting in a maximum global population of less (Craig 2010). While a few smaller colonies have declined or than 47,000 pairs. This lower population estimate cannot been extirpated, the vast majority of colonies lack data that be attributed to a population decline and the data are not would allow assessment of population trends. However, on accurate enough to enable trends to be determined – Green I., near Stewart I./Rakiura, which was reported to quantitative information about population sizes has only have a population of more than a million pairs in the 1940s been collated since the 1990s. Further, there are significant (Stead 1953; Wilson 1959), densities had declined to low challenges in surveying fulmar prions, which – unlike the levels by 2012 (Miskelly 2013c, unpub. data). This is likely other New Zealand nesting prion species – nest mostly in to have been the largest single population in the region, crevices and caves on very remote, seldom-visited islands, and so it can be assumed that the number of fairy prions making them extremely difficult to survey. However, efforts nesting around Stewart I./Rakiura is also likely to be small should be made to monitor at least one colony at each island compared with populations of 70 years ago. group where the species occurs in numbers, e.g. The Pyramid It is recommended that regular surveys of sites through- (Tarakoikoia) (Chatham Is), Proclamation I. (Bounty Is), out the fairy prion’s range are implemented. These are best Toru Islet (Snares Is/Tini Heke) and Ewing I. (Auckland Is). done at Aorangi I. (Northland), Stephens I. (Takapourewa) While historical and recent counts reveal several (Marlborough Sounds), North Brother I. (Marlborough Sounds), Wall I. (West Coast), Motunau I. (Canterbury), significant changes in numbers at prion colonies, the Late Wharekakahu I. (Otago), Tunnel Beach (Otago), Mangere Pleistocene/Holocene fossil record reveals some longer-term I. (Chatham Is), North East I. (Snares Is/Tini Heke) and changes for fairy and broad-billed prions. There is evidence Antipodes I. These islands provide geographic coverage of prehistoric fairy prion colonies on the mainland of New throughout the range of colonies where quantitative data Zealand, with abundant fossils at some South I. sites exist. Ideally, at least two populations in the Stewart I./ (Holdaway et al. 2001). All except a handful of birds nesting Rakiura region should be monitored (e.g. Kundy I. and on the coast of Otago (Loh 2000) have been extirpated, Herekopare I. (Te Marama)) as this region holds the most presumably by introduced mammalian predators. While important fairy prion populations for the southern part of there is evidence for broad-billed prions formerly breeding the species’ New Zealand range. These are both traditional on mainland South I., there is a suggestion that the species muttonbirding islands with restricted access; monitoring may be a recent colonist at the Chatham Is (Tennyson & at these sites could potentially be undertaken by people Millener 1994; Holdaway et al. 2001). with muttonbirding rights on the islands, but would require Our review of published and unpublished data revealed visits outside the March–May muttonbirding season. that very little is known about the population status of any Fulmar prion populations are poorly known. Brooke species of prion nesting in New Zealand, which is particularly (2004) suggests a global population of 50,000–100,000 concerning in light of the loss of 250,000 birds during pairs, with all but the 1000–10,000 pairs estimated to be the 2011 wreck (see above). For each species reviewed, the 76 Tuhinga, Number 27 (2016)

conclusions are similar – there is a lack of reliable data on bycatch in longline fisheries. Endangered Species Research which to assess the population size, status or trends. There is 14: 91–106. a need for population monitoring, even at large colonies, to Anonymous. (1953). Annual report 1939–1940. Pp.3–33. In: Reports and bulletins (1939–1942) of the Ornithological ensure that further localised extinctions are not occurring. Society of New Zealand. xii+98pp. We strongly encourage baseline data to be collected at all Archey, G. and Lindsay, C. (1924). Notes on the birds of major colonies, and the initiation of regular monitoring the Chatham Islands. Records of the Canterbury Museum 2: programmes for all species. This is particularly important 187–201. for fulmar prions due to their small population size and sub- Bartle, J.A. and Paulin, C.D. (1986). Bird observations, Auckland Islands, December 1976. Pp.51–61. In: Penniket, specific diversity (Tennyson & Bartle 2005). We recommend A., Garrick, A. and Breese, E. (compilers). Preliminary reports that surveys employ the methodologies promoted by the of expeditions to the Auckland Islands Nature Reserve 1973– Population and Conservation Status Working Group of 1984. Wellington: Department of Lands and Survey Reserve the Agreement on the Conservation of Albatrosses and Series. 231pp. Petrels (Wolfaardt & Phillips 2013). In brief, we consider Beach, G.S., Wilson, K.-J. and Bannock, C.A. (1997). A survey of the birds, lizards and mammals of Motunau Island, that delineating permanent quadrats at representative Canterbury, New Zealand. Lincoln University Wildlife habitat types throughout key colonies is necessary. Repeated Management Report 14. 19pp. estimates are needed every 5–10 years, counting all burrows Begg, A.C. and Begg, N.C. (1968). Dusky Bay. Christchurch: within each quadrat and checking each burrow’s occupancy. Whitcombe & Tombs. 240pp. These data will allow for the calculation of habitat- Bell, B.D. (1975). Report on the birds of the Auckland Islands Expedition 1972–73. Pp.136–142. In: Yaldwyn, J.C. specific density estimates, which can then be used to assess (compiler). Preliminary results of the Auckland Islands population trends and determine conservation statuses. Expedition 1972–1973. Wellington: Department of Lands and Survey Reserves Series. 447pp. Bell, L.C. (1955). Notes on the birds of the Chatham Islands. Acknowledgements Notornis 6: 65–68. We thank Brian Bell for sharing his unpublished records and Bishop, L.J. (1963). Partial confirmation of two of Reischek’s Little Barrier records. Notornis 10: 306. for his helpful discussion on the state of information; Robin Blackburn, A. (1965). Muttonbird islands diary. Notornis 12: Blyth for her work extracting data from New Zealand 191–207. archives; Matt Charteris for contributions of unpublished Blackburn, A. (1968). The birdlife of Codfish Island. Notornis data; Christine Kiddey (Te Papa) for locating references; and 15: 51–65. Jennifer Twist (Te Papa) for assisting with locating archives. Booth, D.F. (compiler) (1982). Classified summarised notes 30 Data collation was assisted by funding from the Department June 1980 to 30 June 1981. Notornis 29: 49–74. Booth, D.F. (compiler) (1983). Classified summarised notes 30 of Conservation’s Terrestrial and Freshwater Biodiversity June 1981 to 30 June 1982. Notornis 30: 34–68. Information System (TFBIS) fund, while other funding for Brooke, M. de L. (2004). Albatrosses and petrels across the world. the project was contributed by Te Papa. We thank the many Oxford: Oxford University Press. 499pp. people who helped gather data in the field and contributed Brothers, N.P. (1984). Breeding, distribution and status of data to enable this review to be undertaken. We also thank burrow-nesting petrels at Macquarie Island. Australian Wildlife Research 11: 113–131. 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